The Effects of Management Practices on Grassland Birds—Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Chapter PP of

The Effects of Management Practices on Grassland Birds

Professional Paper 1842–PP

U.S. Department of the InteriorU.S. Geological Survey

A

B

C

Cover. A, Female Brown-headed Cowbird. Photograph by David O. Lambeth, used with permission. B, Parasitized Clay-colored Sparrow nest with two Clay-colored Sparrow eggs and three Brown-headed Cowbird eggs. Photograph by Lawrence D. Igl, U.S. Geological Survey. C, Male Brown-headed Cowbird. Photograph by David O. Lambeth, used with permission. Background photograph: Northern mixed-grass prairie in North Dakota, by Rick Bohn, used with permission.

The Effects of Management Practices on Grassland Birds—Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

By Jill A. Shaffer,1 Lawrence D. Igl,1 and Douglas H. Johnson1

Chapter PP ofThe Effects of Management Practices on Grassland BirdsEdited by Douglas H. Johnson,1 Lawrence D. Igl,1 Jill A. Shaffer,1 and John P. DeLong1,2

1U.S. Geological Survey. 2University of Nebraska-Lincoln (current).

Professional Paper 1842–PP

U.S. Department of the InteriorU.S. Geological Survey

U.S. Department of the InteriorDAVID BERNHARDT, Secretary

U.S. Geological SurveyJames F. Reilly II, Director

U.S. Geological Survey, Reston, Virginia: 2019

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Suggested citation:Shaffer, J.A., Igl, L.D., and Johnson, D.H., 2019, The effects of management practices on grassland birds—Rates of Brown-headed Cowbird (Molothrus ater ) parasitism in nests of North American grassland birds, chap. PP of Johnson, D.H., Igl, L.D., Shaffer, J.A., and DeLong, J.P., eds., The effects of management practices on grassland birds: U.S. Geological Survey Professional Paper 1842, 24 p., https://doi.org/10.3133/pp1842PP.

ISSN 2330-7102 (online)

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ContentsAcknowledgments ........................................................................................................................................ivGeneral Overview ..........................................................................................................................................1Rates of Cowbird Parasitism ........................................................................................................................1Host Defenses against Parasitism ..............................................................................................................3Summary..........................................................................................................................................................4References ......................................................................................................................................................4

Figure

PP1. Map showing breeding distribution of the Brown-headed Cowbird (Molothrus ater ) in the United States and southern Canada, based on North American Breeding Bird Survey data, 2008–12 ....................................................................................................................2

Table

PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater ) .......................................................................14

AbbreviationsBBS Breeding Bird Survey

spp. species (applies to two or more species within the genus)

iv

Acknowledgments

Major funding for this effort was provided by the Prairie Pothole Joint Venture, the U.S. Fish and Wildlife Service, and the U.S. Geological Survey. Additional funding was provided by the U.S. Forest Service, The Nature Conservancy, and the Plains and Prairie Potholes Landscape Conservation Cooperative. We thank the following cooperators who provided access to their bibliographic files: Louis B. Best, Carl E. Bock, Brenda C. Dale, Stephen K. Davis, James J. Dinsmore, James K. Herkert, Fritz L. Knopf (deceased), Rolf R. Koford, David R. C. Prescott, Mark R. Ryan, David W. Sample, David A. Swanson, Peter D. Vickery (deceased), and John L. Zimmerman. We thank Anna D. Chalfoun, Stephen K. Davis, Ken D. De Smet, Aaron L. Holmes, Scott D. Hull, Nancy A. Mahony, D. Glen McMaster, Rosalind B. Renfrew, Glenn C. Sutter, and Maiken Winter for sharing information on parasitism rates and unpublished results from their studies. We thank Christopher M. Goldade for his illustration of the Brown-headed Cowbird and the U.S. Geological Survey’s Patuxent Wildlife Research Center, Laurel, Maryland, for providing the range map. We thank Rachel M. Bush, Shay F. Erickson, Susana Rios, and Robert O. Woodward for their assistance with various aspects of this effort. Earlier versions of this chapter benefitted from insightful comments from Mark Fisher and Brian A. Tangen.

The Effects of Management Practices on Grassland Birds—Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

By Jill A. Shaffer, Lawrence D. Igl, and Douglas H. Johnson

General OverviewThe Brown-headed Cowbird (Molothrus ater) is an

obligate brood parasite that relies entirely on host species to incubate its eggs and rear its young. The cowbird evolved on the open grasslands of the Great Plains, but its population expanded eastward and westward following European settle-ment (fig. PP1; Mayfield, 1965; Rothstein and others, 1980; Lowther, 1993). The species’ current breeding distribution is widespread, covering much of the 48 conterminous United States, southern Canada, and northern Mexico. Note that northern Mexico is not shown in figure PP1 because the ability of the North American Breeding Bird Survey to document the status of the Brown-headed Cowbird and other North Ameri-can bird species whose breeding ranges extend into northern Mexico has been limited by a lack of information from poorly surveyed regions in Mexico (Pardieck, 2008). At the conti-nental level, the cowbird reaches its highest abundance in the center of its breeding range in the northern Great Plains, and cowbird abundance generally declines with increasing distance from the Great Plains (Thompson and others, 2000; Igl and Johnson, 2007; Sauer and others, 2013). Vernacular and scien-tific names of plants and animals follow the Integrated Taxo-nomic Information System (https://www.itis.gov/) and the 59th Supplement to the American Ornithological Society’s Check-list of North American Birds (Chesser and others, 2018).

Although cowbirds are not considered a serious threat to their primary avian hosts (Ortega and others, 2005), cowbird brood parasitism has been recognized as a potential threat to the survival of threatened and endangered species (for example, Kirtland’s Warbler [Setophaga kirtlandii]; Kelly and DeCapita, 1982) and has been implicated in the declines of other bird species through reduced reproductive output (Rothstein, 1975, 1990; Mayfield, 1977; Payne, 1977; Brittingham and Temple, 1983; Sealy, 1992; Robinson and others, 1995). As a result, efforts to control or limit cowbird populations in North America are common (Rothstein and Robinson, 1998; Ortega and others, 2005). Elsewhere, Shaffer and others (2003) summarized factors that affect cowbird distribution and parasitic activity and described how cowbirds respond to grassland management practices (for example,

burning, haying, and grazing), including those practices aimed at controlling or limiting cowbird populations. Keys to discouraging Brown-headed Cowbird parasitism or control-ling populations of Brown-headed Cowbirds in the Great Plains are maintaining large expanses of grassland, eliminat-ing foraging areas (for example, feedlots) and perch sites, and reducing the extent of overgrazed pastures (Shaffer and others, 2003). Herein, cowbird parasitism rates are summarized for the 40 species of grassland birds included in “The Effects of Management Practices on Grassland Birds” (Johnson and others, 2019).

Rates of Cowbird ParasitismRates of cowbird brood parasitism, obtained from the

literature and from unpublished data provided by colleagues, are summarized for 40 grassland bird species in table PP1. The frequency of nests parasitized was used as an index to the rates of parasitism, and the rate (percent) of cowbird parasitism was defined as the total number of parasitized nests (×100) divided

Brown-headed Cowbird. Illustration by Christopher M. Goldade, U.S. Geological Survey.

2 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

by the total number of nests reported for a given species in a study.

The Brown-headed Cowbird is a host generalist; eggs of the cowbird have been found in nests of more than 220 species of North American birds (Ortega, 1998; Lowther, 2016). Of the 40 focal grassland bird species, 25 are known victims of cowbird parasitism (table PP1), including four of the 18 nonpasserine species (Greater Prairie-Chicken [Tympa-nuchus cupido], Upland Sandpiper [Bartramia longicauda], Wilson’s Phalarope [Phalaropus tricolor], and Ferruginous

Hawk [Buteo regalis]). Of the 22 grassland passerines, 21 were identified as hosts of the Brown-headed Cowbird (table PP1). There are no known records of cowbird brood parasitism for one passerine, the Sedge Wren (Cistothorus platensis), perhaps because the entrance to its globular-shaped nest is too small for female cowbirds to enter (Herkert and others, 2001). The McCown’s Longspur (Rhynchophanes mccownii) is considered a host based on historical accounts consisting of one to two nests that were parasitized (Fried-mann, 1963), but in records consisting of 10 or more nests,

40°60°80°100°120°140°

50°

30°

10°

Modified from Sauer and others (2014), used with permission from John R. Sauer, U.S. Geological Survey

Base map modified from Esri digital data, 1:40,000,000, 2006Base map image is the intellectual property of Esri and is usedherein under license. Copyright © 2017 Esri and its licensors.All rights reserved.Albers Equal-Area Conic projectionStandard parallels 29°30’ N. and 45°30’ N.Central meridian –96°00’ W.North American Datum of 1983

EXPLANATION

Average number of individuals detected per North American Breeding Bird Survey route per year [>, greater than]

>100

>30 to 100

>10 to 30

>3 to 10

>1 to 3

0.05 to 1

None counted

Not sampled

0 500250 KILOMETERS

0 500250 MILES

Figure PP1. Breeding distribution of the Brown-headed Cowbird (Molothrus ater) in the United States and southern Canada, based on North American Breeding Bird Survey (BBS) data, 2008–12. The BBS abundance map provides only an approximation of breeding range edges.

Host Defenses against Parasitism 3

parasitism was not reported (table PP1). Wilson’s Phalarope and Loggerhead Shrike (Lanius ludovicianus) also exhibited very low parasitism; each species had at least one report of 1 percent (table PP1). The highest parasitism rate was for the Dickcissel (Spiza americana), at 95 percent. Fourteen species had one or more studies reporting heavy (greater than 30 percent) parasitism (as defined by Mayfield, 1965), whereas five species experienced low (less than 10 percent) parasit-ism rates (table PP1). Three species—Sprague’s Pipit (Anthus spragueii), Chestnut-collared Longspur (Calcarius ornatus), and LeConte’s Sparrow (Ammospiza leconteii)—experienced low-to-moderate (10–30 percent) parasitism rates. The remain-ing three parasitized species (Greater Prairie-Chicken, Ferrugi-nous Hawk, and Nelson’s Sparrow [Ammospiza nelsoni]), two of which are nonpasserines, had only one or two incidental records of cowbird parasitism, so a parasitism rate could not be calculated.

Nonpasserine species are generally considered to be unsuitable or inappropriate hosts of the Brown-headed Cowbird because the cowbird’s diet or its developmental strategy (altricial) are incompatible with those of the hosts (Sealy, 2015), and thus, nonpasserine nests usually are avoided by Brown-headed Cowbirds (Rothstein and Robinson, 1998; Sealy, 2015). Sealy (2015) summarized Friedmann’s (1929, 1963) proposed requirements for the suitability of hosts of parasitic cowbirds (Molothrus species [spp.]): (1) the hosts exhibit an altricial post-hatching pattern of development, (2) the hosts lay eggs not much bigger than the cowbird’s eggs, (3) the hosts place food into the nestlings’ gaping bills, and (4) the hosts feed the young an arthropod diet that will sustain the cowbird’s development. None of the four parasit-ized species of nonpasserine grassland birds are known to have raised cowbird young (Ortega, 1998; Sealy, 2015). In contrast, most of the 21 passerine cowbird hosts are known to have fledged cowbird young from parasitized nests (Ortega, 1998; Shane, 2000; Green and others, 2002; Davis and Lanyon, 2008; Davis and others, 2014). Only three grassland passerine hosts are not known to have raised cowbird young: Loggerhead Shrike, McCown’s Longspur, and Nelson’s Sparrow (Ortega, 1998; Yosef, 1996; With, 2010; Shriver and others, 2011).

The reported rates of brood parasitism by Brown-headed Cowbirds vary considerably among and within grassland host species (table PP1), making it difficult to identify generaliz-able patterns across a species’ breeding range. Jensen and Cully (2005a, 2005b) and Igl and Johnson (2007) found evidence of decreasing parasitism levels among grassland birds as distance increases from the center of the cowbird’s breeding range in the Great Plains. A few species with wide-spread breeding distributions follow this pattern, although not consistently, with generally higher parasitism rates in the southern or northern Great Plains and lower parasitism rates east and west of this area. These species include the Savannah Sparrow (Passerculus sandwichensis), Grasshop-per Sparrow (Ammodramus savannarum), Eastern Meadow-lark (Sturnella magna), Bobolink (Dolichonyx oryzivorus),

and Dickcissel (table PP1). Species with breeding ranges restricted to the northern Great Plains that do not conform to this pattern include the Sprague’s Pipit, Chestnut-collared Longspur, McCown’s Longspur, and Baird’s Sparrow (Cent-ronyx bairdii). Of these species, the Baird’s Sparrow was the only species to experience heavy parasitism (36 percent), and that only occurred in one study (Davis, 1994; Davis and Sealy, 1998). The data presented in table PP1 emphasize the complexity and variability of cowbird parasitism in North America and can serve as a reminder that cowbird parasitism rates are not constant across space or time. Brown-headed Cowbirds differentially parasitize hosts within the same habitats, even within the core of the cowbird’s breeding range (Igl and Johnson, 2007). Many factors affect parasitism rates, including location, habitat, proximity of nests to woodland and other habitat edges, alternate hosts, and defensive host behaviors.

Host Defenses against ParasitismMayfield (1965) indicated that grassland birds may have

evolved behavioral defenses against Brown-headed Cowbird brood parasitism because of their long evolutionary history of co-occurrence in the Great Plains. Evolutionarily, however, cowbirds and their hosts have interacted for a relatively short period of time (2.8–3.8 million years) compared to other brood parasite-host systems on other continents (Rothstein and others, 2002; Peer and others, 2013). As such, cowbirds lack some of the more sophisticated adaptations (for example, egg mimicry) possessed by other brood parasites (for example, Common Cuckoo [Cuculus canorus]), and host defenses against cowbird brood parasitism are less complex than defenses exhibited by hosts of other brood parasites (Peer and others, 2013). Defenses possessed by cowbird hosts in grasslands include aggressively defending their nests against female cowbirds, egg rejection, desertion of parasitized nests, and burying cowbird eggs in the nest lining. Many cowbird hosts display aggressive behavior toward cowbirds (Ortega, 1998), including several grassland bird species: Loggerhead Shrike, Clay-colored Sparrow (Spizella pallida), Field Sparrow (Spizella pusilla), Bobolink, and Eastern Meadowlark (Robertson and Norman, 1976, 1977; Grieef, 1995; Ortega, 1998). The success of aggressive behaviors in preventing cowbird parasitism is considered limited (Sealy and others, 1998; Peer and others, 2013). About 13 percent of cowbird hosts are known to reject cowbird eggs (Peer and Sealy, 2004). Of the grassland birds that have been identi-fied as cowbird hosts, the Loggerhead Shrike is considered an egg rejecter (Rothstein, 1982). Several other grassland birds have been recently identified as intermediate rejecters, including the Eastern Meadowlark, Western Meadowlark (Sturnella neglecta), and Dickcissel (Peer and others, 2000). Nest desertion is considered a more effective anti-parasitism strategy. Among smaller cowbird hosts, nest desertion is a

4 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

common response to cowbird parasitism (Ortega, 1998; Hosoi and Rothstein, 2000). Nest desertion with subsequent renest-ing has been reported in about 16 percent of known cowbird hosts (Hosoi and Rothstein, 2000), although the frequency of abandonment may be quite low (Sealy, 1999; Davis and Sealy, 2000). The frequency of desertion of parasitized nests appears to be more common in grasslands and other nonforest habitats than in forest habitats (Hosoi and Rothstein, 2000). Grassland birds known to abandon their nests as a result of parasitism or cowbird removal of host eggs include the Clay-colored Sparrow, Brewer’s Sparrow (Spizella breweri), Field Sparrow, Lark Sparrow (Chondestes grammacus), Lark Bunting (Calamospiza melanocorys), Grasshopper Sparrow, Eastern Meadowlark, and Dickcissel (Baepler, 1968; Hill, 1976; Walley, 1985; Ortega, 1991; Hosoi and Rothstein, 2000). Burial of cowbird eggs by a host is a less common defense against cowbird brood parasitism and is sometimes considered a continuation of nest building (Rothstein, 1975; Sealy, 1995). The Clay-colored Sparrow occasionally buries cowbird eggs in its nest lining (Igl and Johnson, 2007).

SummaryThe Brown-headed Cowbird is an obligate brood parasite

that relies entirely on host species to incubate its eggs and rear its young. The Brown-headed Cowbird is a host gener-alist; eggs of the cowbird have been found in nests of more than 220 species of North American birds. Although cowbirds are not considered a serious threat to their primary avian hosts, cowbird brood parasitism has been recognized as a potential threat to the survival of threatened and endangered species and has been implicated in the declines of other bird species through reduced reproductive output. In this chapter, we summarized the rates of cowbird brood parasitism for 40 grassland bird species. The reported rates of brood parasit-ism by Brown-headed Cowbirds varied considerably among and within grassland host species, and the factors that influ-ence parasitism rates were described. Defenses possessed by cowbird hosts in grasslands include aggressively defending their nests against female cowbirds, egg rejection, desertion of parasitized nests, and burying cowbird eggs in the nest lining.

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6 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

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12 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

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References 13

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14 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Greater Prairie-Chicken (Tympanuchus cupido)

1 record --. Minnesota Svedarsky, 1979.Upland Sandpiper (Bartramia longicauda)

1 record --. Manitoba Davis and others, 1999.1 record --. North Dakota Higgins, 1971.1 record --. Minnesota J.H. Bowles in Friedmann, 1963.1 record --. Montana Igl and Johnson, 2007.

0 15 Nebraska Kempema, 2007.0 16 Oklahoma Reinking and others, 2009.0 21 Wisconsin R.B. Renfrew, Vermont Center for Ecostudies, Norwich, Vermont,

written commun. [n.d.].0 28 Minnesota, North Dakota M. Winter, WissenLeben e.V., Raisting, Germany, and D.H. Johnson,

unpub. data.<1 189 North Dakota Higgins and Kirsch, 1975.

5 43 Nebraska Faanes and Lingle, 1995.8 13 South Dakota Berman, 2007.

Wilson’s Phalarope (Phalaropus tricolor )

1 record --. Manitoba Hatch, 1971.2 records --. Utah Williams and Trowbridge, 1938.

0 21 Minnesota, North Dakota M. Winter and D.H. Johnson, unpub. data.1 386 Saskatchewan M.A. Colwell and D. Delehanty in Colwell and Jehl, 1994.

Ferruginous Hawk (Buteo regalis)

1 record -- North Dakota A. Eastgate in Friedmann, 1929.Loggerhead Shrikeb,c (Lanius ludovicianus)

0 10 Washington Vander Haegen and Walker, 1999.0 60 Missouri Kridelbaugh, 1983.0 61 Minnesota Brooks and Temple, 1990.0 77 Colorado Porter and others, 1975.0 131 Oklahoma Tyler, 1992.0 162 Idaho Woods and Cade, 1996.

<1 1,661 Manitoba K.D. De Smet, Manitoba Conservation Data Centre, Winnipeg, Mani-toba, written commun. [n.d.].

1 261 Iowa De Geus and Best, 1991.Horned Lark (Eremophila alpestris)

0 12 Washington Vander Haegen and Walker, 1999.0 23 Saskatchewan Gaudet, 2013.0 140 Wyoming Mahoney and Chalfoun, 2016.0 163 Saskatchewan Maher, 1973.2 201 Quebec Terrill, 1961.3 161 Ontario Peck and James, 1987.4 119 Ontario Ontario nest records, in Friedmann and others, 1977.

Table PP1 15

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented; --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Horned Lark (Eremophila alpestris)—Continued

9 11 Illinois Robinson and others, 2000.16 32 Minnesota, Montana, North

Dakota, South DakotaIgl and Johnson, 2007.

19 16 Kansas, Missouri, Nebraska Lowther, 1977.19 26 North Dakota R.E. Stewart in Friedmann and others, 1977.45 31 Kansas Hill, 1976.53 15 Saskatchewan S.K. Davis, Canadian Wildlife Service, Regina, Saskatchewan, written

commun. [n.d.].60 84 North Dakota Koford and others, 2000.

Sprague’s Pipit (Anthus spragueii )

0 11 Saskatchewan Lusk, 2009.0 11 Alberta Yoo and Koper, 2017.0 21 Alberta Ludlow and others, 2014.0 24 Saskatchewan B. Dale, unpub. data, in Robbins and Dale, 1999.0 33 Saskatchewan Maher, 1973.0 33 Saskatchewan Gaudet, 2013.0 50 Saskatchewan G.C. Sutter, Royal Saskatchewan Museum, Regina, Saskatchewan writ-

ten commun. [n.d.].0 61 Saskatchewan Pipher, 2011.2 58 Alberta, Saskatchewan Sutter and others, 2016.2 128 Montana Jones and others, 2010.

15 20 Manitoba De Smet, 1992.15 65 Saskatchewan Davis, 2003.16 19 Saskatchewan Klippenstine and Sealy, 2008.18 17 Manitoba Davis, 1994, Davis and Sealy, 2000.

Chestnut-collared Longspur (Calcarius ornatus)

0 36 Saskatchewan Fairfield, 1968.0 38 Saskatchewan Regina Museum of Natural History Nest Record Cards, in Fairfield,

1968.0 54 Saskatchewan Lusk, 2009.0 111 Saskatchewan Maher, 1973.0 115 Saskatchewan Pipher, 2011.0 254 Alberta Hill and Gould, 1997.

<1 212 Saskatchewan Gaudet, 2013.1 137 Alberta Yoo and Koper, 2017.2 770 Montana Jones and others, 2010.4 23 Manitoba Harris, 1944.4 27 Saskatchewan Smith and Smith, 1966.4 71 North Dakota Granfors and others, 2001.5 96 Saskatchewan Klippenstine and Sealy, 2008.7 42 South Dakota Berman, 2007.

16 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Chestnut-collared Longspur (Calcarius ornatus)—Continued8 26 Montana, North Dakota,

South DakotaIgl and Johnson, 2007.

12 26 Manitoba De Smet, 1992.14 57 Manitoba Davis, 1994; Davis and Sealy, 2000.14 352 Montana Lloyd, 2003.16 490 Saskatchewan Davis, 2003.18 22 Saskatchewan Prairie Nest Records Scheme, in Maher, 1973.22 37 North Dakota R.E. Stewart in Friedmann and others, 1977.23 62 North Dakota Stewart, 1975.

McCown’s Longspur (Rhynchophanes mccownii )

1 record --. Saskatchewan Macoun and Macoun, 1909.1 record --. Saskatchewan S.J. Darcus in Friedmann, 1963.1 record --. North Dakota A. Eastgate in Friedmann, 1963.2 records --. North Dakota L.B. Bishop in Friedmann, 1963.

0 45 Wyoming Mickey, 1943.0 60 Montana DuBois, 1935.0 71 Wyoming Mahoney and Chalfoun, 2016.0 74 Saskatchewan Maher, 1973.

Clay-colored Sparrowb,d,e (Spizella pallida)

0 13 Wisconsin Vos and Ribic, 2013.2 250 North Dakota Grant and others, 2006.5 781 Minnesota, North Dakota Winter and others, 2004.9 434 North Dakota Murphy and others, 2017.

10 135 Minnesota Johnson and Temple, 1990.11 204 Manitoba Hill and Sealy, 1994.11 246 North Dakota Pietz and others, 2009.12 196 Minnesota, Montana, North

Dakota, South DakotaIgl and Johnson, 2007.

14 492 North Dakota Granfors and others, 2001.17 496 North Dakota Kerns and others, 2010.20 20 Alberta Salt, 1966.23 13 Ontario Ontario nest records, in Friedmann and others, 1977.24 49 North Dakota Koford and others, 2000.24 275 Alberta, Manitoba, Saskatch-

ewanPrairie Nest Records Scheme, in Friedmann and others, 1977.

27 82 Saskatchewan Davis, 2003.33 40 Minnesota Buech, 1982.36 232 Manitoba Knapton, 1978.38 29 Ontario Peck and James, 1987.38 42 North Dakota Romig and Crawford, 1995.39 33 North Dakota Stewart, 1975.

Table PP1 17

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Brewer’s Sparrowd (Spizella breweri )

0 12 Nevada Rotenberry and Wiens, 1989.0 17 Wyoming A. Chalfoun, U.S. Geological Survey, Laramie, Wyoming, written com-

mun. [n.d.].0 18 Montana L.D. Igl and D.H. Johnson, unpub. data.0 44 Montana Logan, 2001.0 79 Montana B. Walker, Colorado Parks and Wildlife, Grand Junction, Colorado, writ-

ten commun. [n.d.].0 110 Oregon Rotenberry and Wiens, 1989.0 133 Montana A. Chalfoun, written commun. [n.d.].0 153 Wyoming A. Holmes, Northwest Wildlife Science, LLC, written commun. [n.d.].1 428 Montana Abe, 2007.3 32 British Columbia Cannings and others, 1987.4 657 British Colombia Mahony and others, 2006.5 281 Washington Vander Haegen and Walker, 1999.6 363 Washington Vander Haegen, 2007.

13 16 Idaho Rich, 1978.52 25 Alberta Biermann and others, 1987.58 19 Washington N. Mahony, Environment and Climate Change Canada, Edmonton,

Alberta, written commun. [n.d.].Field Sparrowb,d (Spizella pusilla)

<1 681 Pennsylvania Burhans and others, 2001.1 276 Kentucky, Tennessee Giocomo and others, 2008.3 32 Michigan Evans, 1976.3 36 Minnesota, North Dakota M. Winter and D.H. Johnson, unpub. data.3 66 Indiana Nolan, 1963.6 93 Indiana Galligan and others, 2006.7 122 California, Oklahoma, Texas Western Foundation Collection, in Friedmann and others, 1977.

11 36 Michigan Batts, 1958.11 147 Illinois Best, 1978.11 443 Missouri Burhans and others, 2000.13 122 Kentucky, Tennessee Hinnebusch, 2008.14 22 Wisconsin Vos and Ribic, 2013.14 86 Illinois Robinson and others, 2000.16 57 Pennsylvania Norris, 1947.18 33 Michigan Berger, 1951.19 334 Ontario Peck and James, 1987.20 25 Illinois Best, 1979.20 179 Ontario Ontario nest records, in Friedmann and others, 1977.21 52 Michigan Sutton, 1960.27 667 Michigan Walkinshaw, 1978.30 10 Minnesota Buech, 1982.32 72 Michigan Evans, 1978.

18 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Field Sparrowb,d (Spizella pusilla)—Continued32 159 Ohio Hicks, 1934.36 14 Kansas, Missouri, Nebraska Lowther, 1977.49 141 Kansas Rivers and others, 2010.52 71 Illinois Strausberger and Burhans, 2001.53 19 Illinois Strausberger and Ashley, 1997.59 29 Illinois Strausberger, 1998.80 20 Iowa Crooks, 1948; Crooks and Hendrickson, 1953.

Vesper Sparrow (Pooecetes gramineus)

0 10 Iowa Frawley, 1989.0 12 Saskatchewan Maher, 1973.0 18 Michigan, Pennsylvania,

WisconsinHarrison, 1975.

0 19 Saskatchewan Prairie Nest Records Scheme, in Maher, 1973.0 22 Alberta Yoo and Koper, 2017.0 28 Minnesota, North Dakota M. Winter and D.H. Johnson, unpub. data.0 53 North Dakota Granfors and others, 2001.0 56 Saskatchewan Gaudet, 2013.

<1 246 North Dakota Grant and others, 2006.1 37 Saskatchewan Pipher, 2011.1 85 Michigan F.C. Evans in Berger, 1968.2 53 Alberta Ludlow and others, 2014.2 119 Wisconsin Vos and Ribic, 2013.3 77 Washington Vander Haegen and Walker, 1999.3 107 California, Oklahoma, Texas Western Foundation Collection, in Friedmann and others, 1977.4 74 Quebec Terrill, 1961.4 84 Washington Vander Haegen, 2007.5 42 Alberta, Saskatchewan Sutter and others, 2016.8 112 Quebec Hicks, 1934.

10 89 Michigan Southern and Southern, 1980.11 45 Iowa Rodenhouse and Best, 1983.11 442 Ontario Peck and James, 1987.12 195 Alberta, Manitoba,

SaskatchewanPrairie Nest Records Scheme, in Friedmann and others, 1977.

15 20 Minnesota, Montana, North Dakota, South Dakota

Igl and Johnson, 2007.

15 110 Saskatchewan McMaster and others, 1999.15 242 Ontario Ontario nest records, in Friedmann and others, 1977.16 25 Michigan J.F. Ponshair in Berger, 1968.18 44 Saskatchewan Lusk, 2009.20 40 Saskatchewan Klippenstine and Sealy, 2008.26 94 North Dakota Koford and others, 2000.61 41 Iowa Stallman and Best, 1996.

Table PP1 19

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Lark Sparrowd (Chondestes grammacus)

0 18 Kansas Erickson, 2017.0 24 Washington Vander Haegen and Walker, 1999.5 20 Texas Bennett and others, 2014.6 17 Oklahoma Ely, 1957.7 228 California, Oklahoma, Texas Western Foundation Collection, in Friedmann and others, 1977.8 25 Minnesota, North Dakota M. Winter and D.H. Johnson, unpub. data.

10 318 Texas Long and others, 2012.13 15 Manitoba Walley, 1985.13 23 Oklahoma Patten and others, 2011.18 11 Illinois Robinson and others, 2000.20 15 Alberta, Manitoba, Saskatch-

ewanPrairie Nest Records Scheme, in Friedmann and others, 1977.

23 22 Oklahoma Reinking and others, 2009.27 22 Kansas R.F. Johnston in Friedmann, 1963.41 42 Nebraska Kempema, 2007.45 33 Oklahoma Newman, 1970.71 17 Kansas Rivers and others, 2010.82 11 Kansas Hill, 1976.

Lark Buntingd (Calamospiza melanocorys)

0 30 Kansas Shane, 2000.8 12 Nebraska Kempema, 2007.

10 51 Saskatchewan Lusk, 2009.15 142 Kansas Hill, 1976.19 93 Montana, North Dakota,

South DakotaIgl and Johnson, 2007.

21 77 Kansas Wilson, 1976.28 18 Montana, North Dakota,

South DakotaAllen, 1874.

55 22 Saskatchewan Sealy, 1999.61 23 North Dakota Koford and others, 2000.

Savannah Sparrow (Passerculus sandwichensis)

0 26 Michigan Southern and Southern, 1980.0 30 Saskatchewan Pipher, 2011.0 120 Alberta Yoo and Koper, 2017.1 405 Wisconsin Renfrew and others, 2005.2 54 Michigan Potter, 1974.4 27 Saskatchewan Gaudet, 2013.4 140 Quebec Terrill, 1961.7 14 Saskatchewan Prairie Nest Records Scheme, in Maher, 1973.7 531 Ontario Peck and James, 1987.7 675 Minnesota, North Dakota Winter and others, 2004.

20 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Savannah Sparrow (Passerculus sandwichensis)—Continued

10 82 Minnesota, Montana, North Dakota, South Dakota

Igl and Johnson, 2007.

11 190 Alberta Ludlow and others, 2014.11 542 Wisconsin R.B. Renfrew, written commun. [n.d.].13 240 Ontario Ontario nest records, in Friedmann and others, 1977.13 260 Montana Jones and others, 2010.16 83 North Dakota Pietz and others, 2009.16 111 Alberta, Manitoba, Saskatch-

ewanPrairie Nest Records Scheme, in Friedmann and others, 1977.

20 236 North Dakota Kerns and others, 2010.21 24 Manitoba De Smet, 1992.21 150 North Dakota Granfors and others, 2001.22 27 Saskatchewan Lein, 1968.26 141 North Dakota Murphy and others, 2017.28 47 Saskatchewan Klippenstine and Sealy, 2008.29 83 Saskatchewan Davis, 2003.30 100 Montana Fondell and Ball, 2004.32 31 Manitoba Davis, 1994; Davis and Sealy, 2000.37 46 Minnesota Johnson and Temple, 1990.

Grasshopper Sparrowd (Ammodramus savannarum)

0 23 Missouri Winter, 1998.0 78 Wisconsin Renfrew and others, 2005.0 85 Wisconsin Vos and Ribic, 2013.0 131 Kentucky, Tennessee Giocomo and others, 2008.2 41 Indiana Galligan and others, 2006.2 100 Ohio Price, 1934, in Friedmann, 1963.3 32 Wisconsin Byers and others, 2017.4 102 Wisconsin R.B. Renfrew, written commun. [n.d.].4 123 Montana Jones and others, 2010.6 273 Oklahoma Reinking and others, 2009.7 46 Minnesota Johnson and Temple, 1990.7 274 Oklahoma Patten and others, 2011.8 13 Illinois Robinson and others, 2000.8 39 Minnesota, North Dakota M. Winter and D.H. Johnson, unpub. data.8 74 Ontario Peck and James, 1987.9 62 Iowa Patterson, 1994; Patterson and Best, 1996.

10 21 North Dakota Granfors and others, 2001.20 61 Minnesota, Montana, North

Dakota, South DakotaIgl and Johnson, 2007.

21 85 Nebraska Kempema, 2007.22 18 Kansas Hill, 1976.26 58 North Dakota Koford and others, 2000.

Table PP1 21

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Grasshopper Sparrowd (Ammodramus savannarum)—Continued

27 48 Manitoba Davis, 1994; Davis and Sealy, 2000.27 327 Iowa Hovick and others, 2012.30 40 Manitoba De Smet, 1992.33 18 South Dakota Berman, 2007.34 61 Kansas Jensen, 1999.40 81 Kansas Erickson, 2017.44 339 Kansas, Oklahoma Rahmig and others, 2009.48 46 Kansas Rivers and others, 2010.50 18 Kansas Elliott, 1978.58 12 Kansas Klute, 1994; Klute and others, 1997.

Baird’s Sparrow (Centronyx bairdii )

0 11 Saskatchewan Maher, 1973.0 23 Saskatchewan Gaudet, 2013.0 24 Alberta Yoo and Koper, 2017.0 31 Saskatchewan Pipher, 2011.4 89 Montana Jones and others, 2010.5 39 Saskatchewan Lusk, 2009.6 35 Alberta Ludlow and others, 2014.

15 13 Manitoba De Smet and Conrad, 1991.16 68 Manitoba De Smet, 1992.16 70 Saskatchewan Klippenstine and Sealy, 2008.21 182 Saskatchewan Davis, 2003.27 11 North Dakota Granfors and others, 2001.36 76 Manitoba Davis, 1994; Davis and Sealy, 1998.

Henslow’s Sparrow (Centronyx henslowii )

0 20 Wisconsin Byers and others, 2017.0 21 Indiana Galligan and others, 2006.0 35 Missouri Jaster and others, 2014.0 39 Kentucky, Tennessee Hinnebusch, 2008.1 113 Kentucky, Tennessee Giocomo and others, 2008.1 135 Indiana Crimmins and others, 2016.1 136 Indiana J. Robb and others in Herkert and others, 2002.5 59 Missouri Winter, 1999.8 12 Ontario Peck and James, 1987.8 24 Oklahoma Patten and others, 2011.8 24 Oklahoma Reinking and others, 2009.9 22 Oklahoma D. Reinking in Winter, 1999.

22 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

LeConte’s Sparrow (Ammospiza leconteii)

0 36 Wisconsin T.C. Erdman in Lowther, 2005; Richter, 1969.2 50 Minnesota, North Dakota Winter and others, 2005.

25 12 Minnesota, North Dakota, South Dakota

Igl and Johnson, 2007.

29 14 Minnesota Peabody, 1901.Nelson’s Sparrow (Ammospiza nelsoni )

1 record --. Manitoba J. Lane in Hill, 1968.1 record --. Montana Nordhagen and others, 2005.

Bobolinkb (Dolichonyx oryzivorus)

0 12 Illinois Buxton and Benson, 2015.0 19 Michigan Southern and Southern, 1980.0 20 Wisconsin Martin, 1967.5 184 Ohio Hicks,1934.6 52 Wisconsin Byers and others, 2017.6 136 Ontario Peck and James, 1987.

11 314 Minnesota, North Dakota Winter and others, 2004.12 104 Wisconsin R.B. Renfrew, written commun. [n.d.].18 237 Iowa Fletcher and others, 2006.19 78 Minnesota, Montana, North

Dakota, South DakotaIgl and Johnson, 2007.

25 12 North Dakota Koford and others, 2000.25 68 North Dakota Pietz and others, 2009.31 36 North Dakota Granfors and others, 2001.34 47 Minnesota Johnson and Temple, 1990.40 108 North Dakota Kerns and others, 2010.51 839 Nebraska D. Kim in Renfrew and others, 2015.

Eastern Meadowlarkb,c,d (Sturnella magna)

0 11 Wisconsin Vos and Ribic, 2013.0 27 Ohio S.D. Hull, Wisconsin Department of Natural Resources, Madison, Wis-

consin, written commun. [n.d.].0 20 Missouri Jaster and others, 2014.0 87 Kentucky, Tennessee Giocomo and others, 2008.0 131 Indiana Galligan and others, 2006.0 311 Illinois Walk and others, 2010.2 52 Quebec Terrill, 1961.2 57 Illinois Robinson and others, 2000.2 370 Ontario Peck and James, 1987.4 26 Ontario Ontario nest records, in Friedmann and others, 1977.5 582 Oklahoma Reinking and others, 2009.6 36 Wisconsin Byers and others, 2017.6 244 unspecified region Cornell nest records, in Friedmann and others, 1977.6 631 Oklahoma Patten and others, 2011.

Table PP1 23

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Eastern Meadowlarkb,c,d (Sturnella magna)—Continued

10 47 Missouri Winter, 1998.16 38 Wisconsin Lanyon, 1957.22 49 Kansas Erickson, 2017.22 110 Wisconsin R.B. Renfrew, written commun. [n.d.].25 269 Kansas, Oklahoma Rahmig and others, 2009.27 67 Kansas Rivers and others, 2010.31 69 Kansas Jensen, 1999.49 71 Kansas Granfors, 1992.50 10 Kansas Fleischer, 1986.50 14 Kansas Francq, 1972.70 40 Kansas Elliott, 1978.

Western Meadowlarkc (Sturnella neglecta)

0 17 Alberta Yoo and Koper, 2017.0 19 Saskatchewan Gaudet, 2013.0 36 Washington Vander Haegen and Walker, 1999.4 26 Alberta Ludlow and others, 2014.7 29 Kansas Hill, 1976.7 42 Montana Fondell and Ball, 2004.9 54 Nebraska Kempema, 2007.

11 71 Minnesota, North Dakota M. Winter and D.H. Johnson, unpub. data.15 34 Saskatchewan Klippenstine and Sealy, 2008.16 56 Minnesota, Montana, North

Dakota, South DakotaIgl and Johnson, 2007.

18 76 Minnesota Johnson and Temple, 1990.20 20 Saskatchewan Prairie Nest Records Scheme, in Maher, 1973.22 41 Wisconsin Lanyon, 1957.25 103 Saskatchewan Davis, 2003.26 119 Montana Jones and others, 2010.44 36 South Dakota Berman, 2007.44 65 Manitoba Davis, 1994; Davis and Sealy, 2000.45 320 North Dakota Koford and others, 2000.46 24 Manitoba De Smet, 1992.

Dickcisselc,d (Spiza americana)

0 21 Wisconsin Vos and Ribic, 2013.0 29 Illinois Robinson and others, 2000.0 204 Kentucky, Tennessee Giocomo and others, 2008.1 505 Illinois Walk and others, 2010.2 733 Mississippi Conover and others, 2011.3 86 Texas Lituma and others, 2012.3 143 Texas Steigman, 1993.4 47 Indiana Galligan and others, 2006.

24 Rates of Brown-Headed Cowbird (Molothrus ater) Parasitism in Nests of North American Grassland Birds

Table PP1. Rates of brood parasitism in nests of grassland bird species that are known victims of Brown-headed Cowbirds (Molothrus ater).—Continued

[For each species, entries are ordered by increasing parasitism rate. For commonly parasitized species, only reports of 10 or more nests are included. For rarely parasitized species, reports are included regardless of the number of nests to indicate that parasitism has been documented. --, no data; n.d., no date; <, less than]

Parasitism rate (percent)a

Number of nests

Location Source

Dickcisselc,d (Spiza americana)—Continued

5 22 Texas Fretwell and others, 1974.7 269 Oklahoma Churchwell and others, 2008.8 12 Oklahoma Ely, 1957.8 24 Kansas, Missouri, Nebraska Lowther, 1977.9 241 Missouri Winter, 1999.

16 86 Illinois Buxton and Benson, 2015.17 1,208 Oklahoma Patten and others, 2011.18 40 Missouri Jaster and others, 2014.18 1,107 Oklahoma Reinking and others, 2009.19 200 Iowa Fletcher and others, 2006.21 34 Iowa Frawley, 1989.26 23 Kansas R.F. Johnston in Friedmann, 1963.31 61 Oklahoma Overmire, 1962.33 15 Oklahoma Wiens, 1963.33 39 Iowa Patterson, 1994; Patterson and Best, 1996.45 957 Kansas, Oklahoma Rahmig and others, 2009.50 28 Kansas Hill, 1976.50 477 Kansas, Oklahoma Jensen and Cully, 2005b.53 17 Nebraska Hergenrader, 1962.56 124 Kansas Jensen, 1999; Jensen and Finck, 2004.58 184 Kansas Erickson, 2017.60 186 Kansas Hughes, 1996.65 23 Kansas Fleischer, 1986.68 57 Iowa L.D. Igl, unpub. data.69 620 Kansas Zimmerman, 1983.70 890 Kansas Rivers and others, 2010.78 55 Kansas Zimmerman, 1966.82 34 Kansas Klute, 1994; Klute and others, 1997.83 71 Kansas Sandercock and others, 2008.91 65 Kansas Hatch, 1983.95 19 Kansas Elliott, 1978.

aThe following grassland bird species are not known victims of Brown-headed Cowbird brood parasitism: Greater Sage-Grouse (Centrocercus urophasianus), Lesser Prairie-Chicken (Tympanuchus pallidicinctus), Mountain Plover (Charadrius montanus), Long-billed Curlew (Numenius americanus), Marbled Godwit (Limosa fedoa), Willet (Tringa semipalmata), American Bittern (Botaurus lentiginosus), Northern Harrier (Circus hudsonius), Swainson’s Hawk (Buteo swain-soni), Golden Eagle (Aquila chrysaetos), Burrowing Owl (Athene cunicularia), Short-eared Owl (Asio flammeus), Merlin (Falco columbarius), Prairie Falcon (Falco mexicanus), and Sedge Wren (Cistothorus platensis).

bOccasionally displays aggressive behavior toward cowbirds.cEjects cowbird eggs from parasitized nests in some populations.dOccasionally abandons nests due to cowbird brood parasitism.eOccasionally buries cowbird eggs in nest lining.

For more information about this publication, contact:Director, USGS Northern Prairie Wildlife Research Center 8711 37th Street SoutheastJamestown, ND 58401701–253–5500

For additional information, visit: https://www.usgs.gov/centers/npwrc

Publishing support provided by the Rolla Publishing Service Center

Shaffer and others—Rates of B

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Professional Paper 1842–PPISSN 2330-7102 (online)https://doi.org/10.3133/pp1842PP